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Karyotaxonomy of Myosotis Alpestris Group

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Karyotaxonomy of Myosotis Alpestris Group Preslia 78: 345–352, 2006 345 Karyotaxonomy of Myosotis alpestris group Chromozomové počty a taxonomické poznámky k zástupcům skupiny Myosotis alpestris Jitka Š t ě p á n k o v á Institute of Botany, Academy of Sciences of the Czech Republic, CZ-252 43 Průhonice, e-mail: [email protected] Štěpánková J. (2006): Karyotaxonomy of Myosotis alpestris group. – Preslia 78: 345–352. Chromosome numbers of taxa belonging to the Myosotis alpestris group are provided and/or con- firmed. A chromosome count is reported for the first time for M. olympica. A new ploidy level (2n = 24) was revealed within M. stenophylla for which previously only tetraploid cytotypes are reported. Myosotis stenophylla is identified for the first time from Greece. Previous chromosome counts for M. ambigens, M. alpestris, M. atlantica, M. corsicana, M. lithospermifolia, and M. suaveolens are confirmed based on plants originating from karyologically poorly investigated parts of the distribu- tion areas of this polyploid complex. K e y w o r d s : distribution area, karyogeography, Morocco, Myosotis alpestris group, ploidy level, South Europe, taxonomy, Turkey Introduction Representatives of the Myosotis alpestris group are arcto-alpine species with conspicuously scattered distribution areas. The population system of island differentiation is characteristic of this complex; suitable habitats are relatively small but scattered over a large region. Thus, a typical variation pattern develops as a result of limited gene flow, selection pressure of the environment and genetic drift. Along with morphological variation, extensive variability in ploidy level is found in this group. Several factors acting interactively must be invoked to ex- plain the considerably complicated pattern of variability within the M. alpestris group. The most important ones are: polyploidy, ecological adaptation and geographical isolation. In the Myosotis alpestris group, significant chromosome variation among species occurs. Karyological studies of this group have been published by several authors. Of particular in- terest are reports by Grau (1964), Luque (1992) and Štěpánková (1993a, 1993b, 1996). So far three groups of alpine forget-me-nots can be recognized based on their ploidy level: dip- loids (2n = 24), tetraploids (2n = 48) and hexaploids [2n = 72 (70)]. It is quite clear that the process of polyploidization has greatly contributed to the evolution and speciation within this group. Along with conspicuous karyological variation and geographic discontinuities, the species of this group are well known for their interesting ecological disjunction. They grow on serpentine as well as non-serpentine sites both at alpine and lower altitudes. Hence the M. alpestris group might serve as a model for studies of the origin and consequences of geographic and ecological endemism, polyploid evolution and biogeographic history of the European mountain flora. However, the modes of diversification in the group are still un- clear and no phylogenetic hypotheses are proposed. The results of karyological examination presented here are the first part of the project focussed on the biogeographic and evolution- ary history of the complex, combining molecular phylogeny (data from restriction site 346 Preslia 78: 345–352, 2006 analysis of cpDNA and sequence data of ITS region of rDNA) with phylogenetic informa- tion from non-molecular data sets (morphology, karyology). Material and methods Chromosome numbers were counted in samples from 21 populations collected in the field (Table 1) and cultivated in the experimental garden of the Institute of Botany at Průhonice. Counts were usually made on three different root tips from each of two to five individuals cho- sen from each population. Chromosome numbers were determined at the mitotic metaphase in the somatic cells of root tips. The material was pretreated for two hours with a saturated solu- tion of bromnaphtalen and fixed in a freshly prepared 3 : 1 mixture of ethanol and acetic acid (Carnoy). The squash method and staining by lacto-propionic orceine were used. Voucher specimens are deposited in the herbarium of the Institute of Botany at Průhonice (PRA). Results and discussion Myosotis alpestris F. W. Schmidt A species distributed in almost all high mountain ranges in Europe and having broad eco- logical preferences with respect to the geological substrate. Its diagnostic set of morpho- logical characters is: basal leaves with oblanceolate to obovate blade gradually narrowing into a petiol, hirsute with more or less patent hairs or glabrescent on lower surface, calyx narrowed at the base in fruit, with straight, short, appressed hairs not forming conspicuous white rim at the margin of calyx teeth, with few longer, patent, slightly curved hairs and sometimes with few hooked stiff hairs on the calyx tube, not extending up to the flower/ fruit pedicel (Fig. 1a, b), pedicel usually as long as calyx in fruit. Myosotis alpestris is well known for its ability to form cytotypes of different ploidy lev- els. However, this variation in ploidy level is not associated with particular morphological characters. According to previously reported chromosome data (e.g. Grau 1964, Štěpán- ková 1993b), diploid and tetraploid cytotypes occur sympatrically in the Pyrenees, Alps and Carpathians. Hexaploids are known only from the Alps in Germany and Austria (Grau 1964, Štěpánková 1993b). Tetraploid chromosome counts based on plants from the Velebit Mts (Croatia), Prokletije Mts (Montenegro and Albania), Mt Kajmakčalan (Greece), and Pindos Mts (Greece) are reported here for the first time. During the revision of the M. alpestris group in Europe, a noteworthy discrepancy in taxonomic evaluation of sympatric diploid/polyploid cytotypes was noticed. While dip- loids, tetraploids, and hexaploids occupying areas of the Alps and the Carpathians are not usually classified at any taxonomic level (e.g. Grau 1964, Grau & Merxmüller 1972), dip- loids and tetraploids from the Pyrenees are usually treated as separate taxonomic units (Grau 1964, Grau & Merxmüller 1972, Blaise et al. 1992). While the tetraploids are attrib- uted to M. alpestris, diploids are treated as M. pyrenaica (= M. alpina Lapeyer., nom. illeg.; Grau 1964, Grau & Merxmüller 1972), M. corsicana subsp. pyrenaeorum (Blaise et al. 1992), or M. alpestris (Luque 1992). Plants collected in the Pyrenees and included in this karyological analysis bear all of the diagnostic characters of M. alpestris, hence the diploid chromosome number of 2n = 24 reported here is attributed to M. alpestris. Štěpánková: Karyotaxonomy of Myosotis alpestris group 347 Table 1. – List of localities and chromosome numbers of the populations ofMyosotis studied. Taxon Voucher 2n Collection locality Coordinates Altitude Collectors number (m a.s.l) M. atlantica 2006/1 24 Morocco: Haute Atlas Mts, 31°13' N 2550 Štěpánková J., Oukaïmeden 07°51' W Kaplan Z., Kirschner J. 2002 M. alpestris 2005/1 24 Spain: Pyréneés-Orientales 42°41' N 2483 Štěpánková J., Mts, Puerta da la Bonaiqua, 00°59' E 2005 Mt Tuc de la Llanca M. alpestris 2005/2 48 Spain: Pyréneés-Orientales 42°36' N 2380 Štěpánková J. Mts, Parc National de Aigües 00°58' E 2005 Tortes, Port de Ratera M. alpestris 2005/3 48 France: Pyréneés-Orientales 42°34' N 2059 Štěpánková J. Mts, Mt Pic Carlit 01°54' E 2005 M. alpestris 2005/4 48 Montenegro: Prokletije Mts, 42°31' N 1750 Štěpánková J. Gusinje, Mt Volušnica 19°46' E 2005 M. alpestris 2005/5 48 Greece: Northern Pindos Mts, 39°40' N 1640 Štěpánková J. Metsovo, Kataras pass 21°21' E 2003 M. alpestris 2005/6 48 Greece: Nidže voras Mts, Mt 40°54' N 1886 Štěpánková J. Kajmakčalan. 21°49' E 2003 M. alpestris 2005/7 48 Croatia: Velebit Mts, Mt 44°46' N 1620 Štěpánková J. Crikvena 14°59' E 2005 M. ambigens 2005/8 24 Italy: Monte Baldo Mts, the 45°38' N 1500 Štěpánková J., crest Creta di Naole 10°47' E Kaplan Z. 2004 M. corsicana 2005/9 24 Corsica: Vizzavona, Mt Monte 42°08' N 2203 Šafránek J. D’Oro 29°06' E 2004 M. lithospermifolia2005/10 24 Turkey: Esler Dagi Mts, Mt 37°40' N 1950 Štěpánková J. Honaz Dag 29°18' E 2004 M. lithospermifolia2005/11 24 Turkey: Dedegöl Daglari Mts, 37°41' N 2036 Štěpánková J. Mt Dededöl Dagi 31°18' E 2004 M. lithospermifolia2005/12 24 Turkey: Ak Daglar Mts: Mt 36°31' N 2108 Štěpánková J. Kofu Tepe 29°55' E 2004 M. olympica 2005/13 24 Turkey: Uludag Mts, Mt Uludag40°05' N 2232 Štěpánková J. Tepe 29°11' E 2004 M. stenophylla 2005/14 24 Greece: Northern Pindos Mts, 40°06' N 2087 Štěpánková J. Mt Smolikas 20°54' E 2003 M. stenophylla 2005/15 24 Greece: Northern Pindos Mts, 40°05' N 2600 Štěpánková J. Mt Smolikas 20°55' E 2003 M. suaveolens 2005/16 24 Greece: Central Olympus Mts, 40°05' N 2815 Štěpánková J. Mt Skolio 22°22' E 2003 M. suaveolens 2005/17 24 Montenegro: Prokletije Mts, 42°30' N 1790 Štěpánková J. Gusinje, Mt Karanfili 19°47' E 2005 M. suaveolens 2005/18 24 Albania: Prokletije Mts, Mt 42°27' N 2308 Štěpánková J. Maja e Jezerce 19°49' E 2005 M. suaveolens 2005/19 24 Montenegro: Komovi Mts: 42°43' N 2100 Štěpánek J., Jr. Berane, Mt Kom Vasojevički 19°41' E 2005 M. suaveolens 2005/20 24 Montenegro: Moračke Planine 42°50' N 1840 Štěpánková J. Mts: Berane, Mt Moračka Kapa 19°15' E 2005 348 Preslia 78: 345–352, 2006 Myosotis ambigens (Bég.) Grau A species considered to be restricted to the Apennine Mts (Italy) (Grau 1964, Grau & Merxmüller 1982). They are small mat-forming plants with small short petiolate basal leaves and narrowly oblanceolate blade hirsute with long patent hairs on both surfaces or sometimes glabrescent on the lower surface, gradually narrowing into a petiole, with con- spicuously smaller linear to lanceolate hirsute cauline leaves (ca 5 × 15 mm in the middle part of stem), obtuse at the apex, with fruiting calyx more or less narrowed at the base, densely covered with short hairs and with numerous long stiff appressed white hairs, forming conspicuous white rim at the margin of calyx teeth, calyx tube with no or very few hooked hairs, pedicel without hooked hairs (Fig.
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